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Sarcodina: Amoebae NOAA Technical Report NMFS Circular 419 Marine Flora and Fauna of the Northeastern United States. Protozoa: Sarcodina: Amoebae Eugene C. Bovee and Thomas K. Sawyer January 1979 U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps, Secretary National Oceanic and Atmospheric Administration Richard A. Frank, Administrator Terry L. Leitzell, Assistant Administrator for Fisheries National Marine Fisheries Service For S;le!:;y the· Superintendent of -DOeum~;:':ts-:-U.S. Government" Printi;:;-g -offict;' Washington, D.C. 20402 Stock No. 003-017-00433-3 FOREWORD This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the Northeastern Unit.ed States." This subseries will consist of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and animals of the northeastern United States. Manuals will be published at irregular intervals on as many taxa of the region as there afe specialists available to collaborate in their preparation. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under the auspices of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. Instead of revising the "Woods Hole Keys," the staff of the Systematics-Ecology Program decided to ex­ pand the geographic coverage and bathymetric range and produce the keys in an entirely new set of expanded publications. The "Marine Flora and Fauna of the ~ortheastern United States" is being prepared in collaboration with systematic specialists in the United States and abroad. Each manual will be based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals. Each major taxon, treated in a separate manual, will include an intro­ duction, illustrated glossary, uniform originally illustrated keys, annotated check list with infor­ mation when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a systematic index. These manuals are intended for use by biology students, biologists, biological oceanographers, informed laymen, and others wishing to identify coastal organisms for this region. In many instances the manuals will serve as a guide to additional information about the species or the group. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is planned to include organisms from the headwaters of estuaries seaward to approximately the 200-m dept.h on the continental shelffrom 1\1aine10 Virginia, but may vary somewhat with each major taxon and the interests of collaborators. Whenever possible representative specimens dealt with in the manuals will be deposited in reference collections of major museums, After a sufficient number of manuals of related taxonomic groups have been published, the manuals will be revised, grouped, and issued as special volumes. These volumes will thus con­ sist of compilations of individual manuals within phyla such as the Coelenterata, Arthropoda, and Mollusca, or of groups of phyla. CONTENTS Page Introduction ..... 1 Diagnostic characters 1 Ecology . 2 Collecting methods 2 Culture . 2 Observational techniques 2 Common problems in determining the identity of marine Sarcodina 3 Glossary . 3 Key to families and genera of Amoebae and descriptions of species 6 Annotated systematic list 49 Selected bibliography 52 Systematic index 54 Acknowledgments .. 55 Coordinating editor's comments 56 The National Marine Fisheries Service (NMFS) does not approve, rec~ ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, Of to this publication furnished by NMFS, in any advertising or sales pro­ motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. iii Marine Flora and Fauna of the Northeastern United States. Protozoa: Sarcodina: Amoebae EUGENE C. BOVEE' and THOMAS K. SAWYER' ABSTRACT This manual contain8 a key to 15 families of freshwater and marine amoebae, ofwhich only one, the Echinamoebidac, docs not contain a known marine species. Diagnostic features for 49 genera, of which 34 include marine species, also are given. Descriptions and illustrations for 76 species ofmarine amoebae and an annotated systematic list are provided. The basic key is designed to assist the user in the identification ofrecognizcd species ofmarine amoebae that have been described from waters of the northeastern United States. However, certain well-known families and genera of freshwater forms are included to assist in thci'r identification should they be discovered in seawater in future investiga­ tions. Information also is provided which includes comments on the general biology ofthe Amoebida, and techniques for microscopic observations and laboratory cultivation of many species. Most of the amoebae described in the key are free living, but a few are parasitic al1d known to be of considerable economic importance. One new free-living species, Vexillifera minutissima, was discovered in Chin­ coteague Bay, Va., and is described herein for the first time. INTRODUCTION Amoebae usually are difficult to preserve in their typical locomotive form. Sketches and photographs The amoebae are that group of Sarcodina (Protozoa) made by dependable taxonomists of amoebae are as yet which move by means of subtle changes in the physical the best sources for their identification, while fixed and properties of their protoplasm. Probable gel-sol inter­ stained specimens are useful for studying their nuclear conversions of their protoplasm produce extensions of the structures. There are no extensive type collections, but a body mass to form pseudopods which are involved in few protozoologists maintain certain species in con­ both locomotion and feeding activities. The movement of tinuous culture. the protoplasm may be cyclic in the body mass and in the Photomicrographs are not an entirely adequate sub­ pseudopods which conduct different specialized func­ stitute for illustration of type-specimens and descrip­ tions. The protoplasm is convertible from a slightly vis­ tions should be supplemented with detailed sketches cous fluid state (plasmasol) to a more viscous or rigid which are representative of the species being described; contractile jelly (plasmagel). The specific morphological photomicrographs show only a planar section through appearance of various Sarcodina is a dynamic product of the amoeba. Scanning electronmicrographs are not yet their protoplasmic activities and movements. available for most amoebae, and, besides being costly to The amoebae of the ocean resemble those of fresh­ prepare, have the disadvantage of portraying fixed water ponds and streams, and the same group of charac­ specimens whose contours are more or less distorted. ters can be used to distinguish them. The amoebae of the Hence, few amoebae have been illustrated with scan­ ocean are not yet well known. A great many unknown ning techniques and those available do not substitute for species remain to be found, described, identified, and direct microscopic identification. classified. However, enough of them are known to Fixed and stained specimens may be prepared by us­ provide a basis for provisional identification to the ing routine cytological methods for staining cells (Alger generic level. Known species are included as examples. 1966; :Mitchell 1966); including such stains as iron­ Since identification of amoebae classically depends hematoxylin, Kernecht's-red, Feulgen's reaction, mainly on their forms and activities while in loco­ chromotrope-2R, or Biebrich's scarlet-fast-green. For motion, observations must be made while they are ac­ electronmicroscopy and light microscopy, various tive. Many marine amoebae are active at dawn or dusk, modifications of combined osmic acid and glutaralde­ particularly the many small species found feeding on hyde fixations have been used with success. bacteria in bays and estuaries. Others are active mainly during midday. Observations at almost any time from DIAGNOSTIC CHARACTERS dawn to dark, or even at night, may turn up an active amoeba. Members of the Sarcodina, usually considered to be a subphylum of the Protozoa, are typically single celled; but some are multinucleate, and others are plasmodial, 'Department of Physiology and Cell Biology'. Division of Biological Sciences. University of Kansas. Lawrence. KS Gl1045. that is, they can fragment into uninucleate cytoplasmic 'Kortheast Fisheries Center. NMFS, NOAA. Oxford. :\1D 21654. parts which may fuse again into a multinucleate cyto- 1 plasmic mass. In most Sarcodina the nucleus is vesic­ amoebae are important organisms in oceanic ecosys­ ular, i.e., round and bounded by its own membrane, and tems. usually contains a round central endosome or nucleolus or both. The cytoplasm is motile within a mucoid outer COLLECTING METHODS coat (called a plasmalemma) and is extensible into lobes or threads, called pseudopods, which are also used in Amoebae may be found in almost any damp locomotive progress and feeding. The plasmalemma or place-moist soil or sand, on aquatic vegetation, on wet outer coat may bear a fuzz-like layer which
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